Phylogeny of Lithobiidae Newport, 1844, with emphasis on the megadiverse genus Lithobius Leach, 1814 (Myriapoda, Chilopoda).

Phylogenetic analyses based on molecular and morphological data were conducted to shed light on relationships within the mostly Palaearctic/Oriental centipede family Lithobiidae, with a particular focus on the Palaearctic genus Lithobius Leach, 1814 (Lithobiidae, Lithobiomorpha), which contains >500 species and subspecies. Previous studies based on morphological data resolved Lithobius as nonmonophyletic, but molecular-based phylogenetic analyses have until now sampled few species. To elucidate species inter-relationships of the genus, test the validity of its classification into subgenera, and infer its relationships with other Lithobiidae, we obtained molecular data (nuclear markers: 18S rRNA, 28S rRNA; mitochondrial markers: 16S rRNA, COI) and 61 morphological characters for 44 species of Lithobius representing four of its eight subgenera and nine other representatives of Lithobiidae. The data were analyzed phylogenetically using maximum-likelihood, parsimony and Bayesian inference. This study suggests that (i) a close relationship between L. giganteus and the pterygotergine Disphaerobius loricatus highlighted in recent morphological analyses is also strongly supported by molecular data, and Pterygoterginae is formally synonymized with Lithobiinae; (ii) the Oriental/Australian genus Australobius is consistently resolved as sister group to all other sampled Lithobiidae by the molecular and combined data; (iii) the subfamily Ethopolyinae may be paraphyletic; (iv) the genus Lithobius is nonmonophyletic; (v) the subgenera Lithobius, Sigibius and Monotarsobius are nonmonophyletic and should not be used in future taxonomic studies; and (vi) there are instances of cryptic species and cases in which subspecies should be elevated to full species status, as identified for some European taxa within Lithobius.

Segmentation of the millipede trunk as suggested by a homeotic mutant with six extra pairs of gonopods.

BACKGROUND: The mismatch between dorsal and ventral trunk features along the millipede trunk was long a subject of controversy, largely resting on alternative interpretations of segmentation. Most models of arthropod segmentation presuppose a strict sequential antero-posterior specification of trunk segments, whereas alternative models involve the early delineation of a limited number of 'primary segments' followed by their sequential stereotypic subdivision into 2n definitive segments. The 'primary segments' should be intended as units identified by molecular markers, rather than as overt morphological entities. Two predictions were suggested to test the plausibility of multiple-duplication models of segmentation: first, a specific pattern of evolvability of segment number in those arthropod clades in which segment number is not fixed (e.g., epimorphic centipedes and millipedes); second, the occurrence of discrete multisegmental patterns due to early, initially contiguous positional markers. RESULTS: We describe a unique case of a homeotic millipede with 6 extra pairs of ectopic gonopods replacing walking legs on rings 8 (leg-pairs 10-11), 15 (leg-pairs 24-25) and 16 (leg-pairs 26-27); we discuss the segmental distribution of these appendages in the framework of alternative models of segmentation and present an interpretation of the origin of the distribution of the additional gonopods.The anterior set of contiguous gonopods (those normally occurring on ring 7 plus the first set of ectopic ones on ring 8) is reiterated by the posterior set (on rings 15-16) after exactly 16 leg positions along the AP body axis. This suggests that a body section including 16 leg pairs could be a module deriving from 4 cycles of regular binary splitting of an embryonic 'primary segment'. CONCLUSIONS: A very likely early determination of the sites of the future metamorphosis of walking legs into gonopods and a segmentation process according to the multiplicative model may provide a detailed explanation for the distribution of the extra gonopods in the homeotic specimen. The hypothesized steps of segmentation are similar in both a normal and the studied homeotic specimen. The difference between them would consist in the size of the embryonic trunk region endowed with a positional marker whose presence will later determine the replacement of walking legs by gonopods.

A new species of Lophostreptus Cook, 1895 discovered among syntypes of L.regularis Attems, 1909 (Diplopoda, Spirostreptida, Spirostreptidae).

A new species of the genus Lophostreptus Cook, 1895 is described, based on specimens hidden for over a century among the syntypes of its congener Lophostreptusregularis Attems, 1909 housed in the Naturhistoriska Riksmuseet Stockholm (NRMS) and the Naturhistorisches Museum Wien (NHMW). A lectotype is designated for Lophostreptusregularis Attems, 1909 in order to stabilize its taxonomy. Updates to the millipede fauna of Mt. Kilimanjaro, Tanzania are provided.

An annotated catalogue of selected historical type specimens, including genetic data, housed in the Natural History Museum Vienna.

Museum collections are an important source for resolving taxonomic issues and species delimitation. Type specimens as name-bearing specimens, traditionally used in morphology-based taxonomy, are, due to the progress in historical DNA methodology, increasingly used in molecular taxonomic studies. Museum collections are subject to constant deterioration and major disasters. The digitisation of collections offers a partial solution to these problems and makes museum collections more accessible to the wider scientific community. The Extended Specimen Approach (ESA) is a method of digitisation that goes beyond the physical specimen to include the historical information stored in the collection. The collections of the Natural History Museum Vienna represent one of the largest non-university research centres in Europe and, due to their size and numerous type specimens, are frequently used for taxonomic studies by visiting and resident scientists. Recently, a version of ESA was presented in the common catalogue of the Fish and Evertebrata Varia collections and extended to include genetic information on type specimens in a case study of a torpedo ray. Here the case study was extended to a heterogeneous selection of historical type series from different collections with the type locality of Vienna. The goal was to apply the ESA, including genetic data on a selected set of type material: three parasitic worms, three myriapods, two insects, twelve fishes, and one bird species. Five hundred digital items (photographs, X-rays, scans) were produced, and genetic analysis was successful in eleven of the 21 type series. In one case a complete mitochondrial genome was assembled, and in another case ten short fragments (100-230 bp) of the cytochrome oxidase I gene were amplified and sequenced. For five type series, genetic analysis confirmed their taxonomic status as previously recognised synonyms, and for one the analysis supported its status as a distinct species. For two species, genetic information was provided for the first time. This catalogue thus demonstrates the usefulness of ESA in providing digitised data of types that can be easily made available to scientists worldwide for further study.

Ommatoiulus ktarii, a new millipede from Tunisia and considerations on the taxonomy of the genus (Diplopoda, Julida, Julidae).

A new millipede, Ommatoiulus ktarii sp. nov., is here described from Tunisia based on relatively recently collected specimens from the semi-arid area of the plain of Oueslatia. A description of the external morphology and gonopod structures are provided, proving the new species to be most similar to the only hitherto known schizophyllinine from Libya, Ommatoiulus rimosus (Karsch, 1881) described more than 140 years ago. Photographs of the gonopods of all Tunisian Ommatoiulus Latzel, 1884 species are provided.

The genus Inversotyphlus Strasser, 1962, stat. nov. and Inversotyphlusammirandus sp. nov., a new bizarre, highly modified troglobiotic millipede (Diplopoda, Julida, Julidae) from Albania, Balkan Peninsula.

Inversotyphlus Strasser, 1962, stat. nov. is raised to the genus level and a new hydrophilous species, Inversotyphlusammirandussp. nov., is described from the second deepest pit in Albania. This species is characterized by a highly modified head and body for a presumably semiaquatic or hygropetric life and filtering diet. It is by far the most bizarrely modified cave-dwelling julid known. The new species is diagnosed, described in detail, and richly illustrated. Besides I.ammirandussp. nov., the genus Inversotyphlusstat. nov. includes six species: I.clavatus (Antic, 2018), comb. nov., I.edentulus (Attems, 1951), comb. nov., I.gellianae (Makarov & Rada, 2006), comb. nov., I.gracilis (Antic, 2018), comb. nov., I.lobifer (Attems, 1951), comb. nov., and I.opisthonodus (Antic, 2018) comb. nov. The subgenus Attemsotyphlussyn. nov. is here considered as a junior subjective synonym of the genus Inversotyphlusstat. nov. Notes are given on the habitat of I.ammirandussp. nov., the taxonomy of the tribe Typhloiulini and the genus Inversotyphlusstat. nov., and adaptive modifications of the mouthparts.

Integrative redescription of the sucking millipede genus Dawydoffia Attems, 1953, with a description of a new species and a transfer to the family Hirudisomatidae (Diplopoda, Polyzoniida).

The type species of the monotypic Polyzoniida genus Dawydoffia Attems, 1953, D. kalonota Attems, 1953 from Vietnam, is redescribed based on type material. A second species of the genus, D. siphonocryptida n. sp., is described from Laos based on scanning electron microscopy, micro-computed tomography and molecular barcoding. The species of Dawydoffia are among the shortest and widest of the Polyzoniida, and resemble in habitus those of the Siphonocryptida genus Siphonocryptus Pocock, 1894, both having pleurites that are completely fused to the tergites. Dawydoffia was previously placed in the family Siphonotidae Cook, 1895, but can be identified as a member of the Holarctic Hirudisomatidae Silvestri, 1896 based on the following morphological characters: the uplifted posterior margins of the tergites, the collum covering part of the head, the position of the male gonapophysis or pseudopenis, the retracted telson, and the ozopores situated close to the tergal margin. However, both Dawydoffia species have a slender paronychium, a character previously known only from the Siphonotidae, but also documented here for Hirudisoma roseum (Victor, 1839). A slight redefinition of the Polyzoniida families is provided. The two Dawydoffia species differ in their coloration, as well as in their somatic and gonopodal characters.

A revision of the millipede family Paracortinidae (Diplopoda, Callipodida).

The taxonomy of the family Paracortinidae Wang & Zhang, 1993 is revised based on literature, old and recently collected material. A new genus Crassipetalum Akkari & Stoev, gen. nov. is described, to accommodate a new species Crassipetalummagnum Akkari & Stoev, gen. nov. et sp. nov. and a recently described species Crassipetaluminflatum (Chen, Zheng & Jian, 2023), comb. nov. The genus Scotopetalum Shear, 2000 hitherto described for the Vietnamese species S.warreni Shear, 2000 and subsequently synonymised with the genus Paracortina Wang & Zhang, 1993 is here resurrected and supplemented with another species, S.chinensis (Stoev & Geoffroy, 2004), comb. nov., ex Paracortinachinensis Stoev & Geoffroy, 2004. The status of the fourth genus in the family, Angulifemur Zhang, 1997, is reconfirmed. Based on recently collected specimens from China, two new species of the genus Paracortina are described: P.asciformis Akkari & Stoev, sp. nov. (Sichuan Prov., Lixian County) and P.kabaki Akkari & Stoev, sp. nov. (Yunnan, Shangrila County). The Vietnamese species Paracortinamultisegmentata Stoev & Geoffroy, 2004 and Paracortinakyrang Nguyen, Stoev, Nguyen & Vu, 2023 are considered of uncertain taxonomic position within Paracortinidae. Differential diagnoses for the proposed genera as well as for all the species are presented, and descriptions or descriptive notes for all the species are provided, using a standardised terminology, and when possible, accompanied by micrographs of the habitus and gonopod structures. An identification key based on gonopod structures is proposed for all the members of the family. A discussion on species affinities, secondary sexual characters, troglomorphic characters, geographical distribution, and habitat preferences are also provided together with a distribution map for all members of the family.

Sexually dimorphic characters of the ultimate legs in lithobiid centipedes (Myriapoda, Chilopoda, and Lithobiomorpha): Morphology and implications for reproductive behavior.

Many species of lithobiomorph centipedes present a pronounced sexual dimorphism reflected in remarkable structural modifications on the ultimate legs of males. Most records of these male secondary sexual characters addressed taxonomy, helping to identify and characterize species or diagnose genera, but information on their diversity, detailed morphology and possible function(s) is scarce. In this study, nine species of the two lithobiid genera Lithobius Leach, 1814 and Eupolybothrus Verhoeff, 1907 were investigated, using light and scanning electron microscopy to document the detailed morphology of secondary sexual characters of male ultimate legs. Secondary sexual characters affecting the cuticle of the ultimate legs are described in detail and found to often be associated with sensilla, interpreted here as chemo- and mechanoreceptors, and with clusters of pores, a hitherto undescribed pore-distribution for this group. The tibial nodule of the species Lithobius nodulipes Latzel, 1880, was additionally examined with histological semi-thin sections. These results revealed that the clustered pores are connected to glandular tissue, and are, based on their morphology, interpreted as openings of flexo-canal epidermal glands. The presence of various sensory and glandular structures associated with sexual dimorphism indicates a likely role during courtship and mating. The closely related species examined in this research show comparable dimorphic structures, which are otherwise species-specific. Morphological observations on secondary sexual structures inform on reproductive biology in groups like lithobiomorphs for which there are limited behavioral data.

A new species of the hitherto monospecific genus Pleonoporus Attems, 1938 (Diplopoda, Spirostreptida, Odontopygidae).

The hitherto monospecific genus Pleonoporus is revised based on the syntypes of P.robustus Attems, 1938, housed in Naturhistorisches Museum Wien (NHMW), as well as on specimens of what we interpret as a new species of the genus, Pleonoporustanzanicus sp. nov., collected in Tanzania and housed in the Museum of Nature - Zoology, Leibnitz Institute for the Analysis of Biodiversity Change (ZMH) for more than a century. Both species are described based on habitus and gonopod structures and illustrated with micrographs, whereas scanning electron microscope images are also provided for the new species. This paper further highlights the importance of natural history collections for taxonomic research and studies on species diversity in general.

A review of the cavernicolous Trichopolydesmidae (Diplopoda, Polydesmida) from the Carpathian-Balkan arch and the Rhodope Mountains, with descriptions of two new genera and three new species.

All cavernicolous species of the millipede family Trichopolydesmidae from the Carpathian-Balkan arch and the Rhodope Mountains have been reviewed. At present the family has been shown to comprise five or six genera with eight or nine species. Two new genera have been described, viz., Balkanodesminus gen. nov., with two new species: B.dentatoides sp. nov. and B.serbicus sp. nov., from Bulgaria and Serbia, respectively, and the monospecific Rhodopodesmus gen. nov., with R.niveus sp. nov., from Bulgaria. Two new combinations and one new status have been proposed: Balkanodesminusbulgaricus (Strasser, 1962) comb. nov. ex Bacillidesmusbulgaricus Strasser, 1962 and Balkanodesminusdentatus (Strasser, 1966a) comb. nov., stat. nov. ex Bacillidesmusbulgaricusdentatus Strasser, 1966a. All genera and species are diagnosed with the inclusion of the most relevant remarks for each of them. Old museum types are checked for Bacillidesmusfiliformis (Latzel, 1884) with lectotype designation, as well as for Trichopolydesmuseremitis Verhoeff, 1898. An identification key to all six genera and a distribution map for the eight species are provided, as well as brief remarks and general considerations on the family Trichopolydesmidae.

Review of Trachysphaera Heller, 1858 (Diplopoda: Glomerida: Glomeridae) in Serbia, with taxonomic notes on the genus.

The glomerid genus Trachysphaera Heller, 1858 in Serbia is revised, based on recently collected specimens and historical material. Six species are presently recorded from Serbia, viz., Trachysphaera corcyraea (Verhoeff, 1900), T. cristangula (Attems, 1943), T. schmidtii Heller, 1858, and T. similicostata (Radu Ceuca, 1951), all new to the fauna of the country, as well as T. costata (Waga, 1857a) and T. lobotarsus (Attems, 1943). All of these species are richly illustrated and mapped. To verify the identity of some species, historical material from the Natural History Museum in Vienna, including syntypes, is also studied, and two lectotypes have been designated. Some misidentifications from the literature have been corrected. Trachysphaera acutula (Latzel, 1884) syn. n., T. cultrifera (Verhoeff, 1906) syn. n. and T. multiclavigera (Verhoeff, 1898) syn. n. are all considered as junior subjective synonyms of T. schmidtii. Trachysphaera attemsi Golovatch, 1976 syn. n. is transferred from the list of synonyms of T. costata to the list of synonyms of T. corcyraea. One species, T. similicostata, is resurrected from synonymy. Relationships between some congeners and the taxonomic problems within the genus are briefly discussed.

Haasea Verhoeff, 1895-a genus of tumultuous history and chaotic records-redefinition, revision of taxonomy and geographic distributions, with descriptions of two new species from Austria and Serbia (Diplopoda, Chordeumatida, Haaseidae).

In this paper we revise the chordeumatidan genus Haasea Verhoeff, 1895 for the first time and describe two new species viz., H. gruberi sp. nov. and H. makarovi sp. nov. discovered in Austria and Serbia, respectively. A new terminology is proposed to standardize the description of the gonopod structures in the genus. Type material obtained from several institutions was examined and documented herein, whereby lectotypes are designated. We clarified the taxonomic status of a few taxa and consider the species H. norica (Verhoeff, 1913) and H. guidononveilleri Makarov, 2008 as a junior subjective synonyms of H. cyanopida (Attems, 1903) and H. vidinensis (Strasser, 1973a), respectively. The following subspecies or varieties, viz., Orobainosoma filicis ossiacum Verhoeff, 1939b, Orobainosoma flavescens vornatscheri Verhoeff, 1935, Orobainosoma hungaricum orientale Tabacaru, 1965, and Orobainosoma inflatum var. aspidiorum Verhoeff, 1929 are here regarded as junior subjective synonyms of their nominal taxa. The taxonomic relationships within the genus Haasea are discussed and an identification key to its 17 species is presented, based on number of body segments and gonopod structures.                Maps showing species occurences based on historical and recent records are presented to understand their respective geographic ranges. H. flavescens is now deleted from the list of species in Croatia, Bosnia and Herzegovina, Slovenia and Italy as these records were based on misidentified material. H. inflata is deleted from the fauna of Bosnia and Herzegovina, whereas H. hungarica is newly recorded from Austria, Serbia and Slovenia.

Five million years in the darkness: A new troglomorphic species of Cryptops Leach, 1814 (Chilopoda, Scolopendromorpha) from Movile Cave, Romania.

A new species of Cryptops Leach, 1814, C. speleorex sp. nov., is described from Movile Cave, Dobrogea, Romania. The cave is remarkable for its unique ecosystem entirely dependent on methane- and sulfur-oxidising bacteria. Until now, the cave was thought to be inhabited by the epigean species C. anomalans, which is widespread in Europe. Despite its resemblance to C. anomalans, the new species is well-defined morphologically and molecularly based on two mitochondrial (cytochrome c oxidase subunit I COI and 16S rDNA) and one nuclear (28S rDNA) markers. Cryptops speleorex sp. nov. shows a number of troglomorphic traits such as a generally large body and elongated appendages and spiracles, higher number of coxal pores and saw teeth on the tibia of the ultimate leg. With this record, the number of endemic species known from the Movile Cave reaches 35, which ranks it as one of the most species-rich caves in the world.

On the identity of the centipede Sogona cyclareata Attems, 1947 (Geophilomorpha, Geophilidae).

The type material of Sogona cyclareata Attems, 1947 is revised. The syntype shares characters with representatives of the family Schendylidae and not the Geophilidae to which it is currently assigned. The species has the claw of the second maxillae pectinate and the labrum not divided into median and lateral portions. Sogona cyclareata presents also characters of the genus Schendylops Cook, 1899 such as the presence of two pores in each coxopleuron, ultimate legs without claws, and the arrangement of the sternal pores-fields. No Schendylops species known to date has the characters of S. cyclareata, so we propose the new combination Schendylops cyclareatus.

A troglobitic species of the centipede Cryptops (Chilopoda, Scolopendromorpha) from northwestern Botswana.

A new species of Cryptops, C. (Cryptops) legagussp. nov., occurs in caves in the Koanaka and Gcwihaba Hills in northwestern Botswana. Bayesian molecular phylogenetics using 18S rRNA, 28S rRNA, 16S rRNA and cytochrome c oxidase subunit I corroborates a morphological assignment to the subgenus Cryptops and closest affinities to southern temperate species in South Africa, Australia and New Zealand. The new species is not conspicuously modified as a troglomorph.

The Myriapoda of Halimun-Salak National Park (Java, Indonesia): overview and faunal composition.

The myriapod fauna of the mega-diverse country of Indonesia is so far insufficiently known, with no species lists or determination keys. In order to obtain an overview of the faunal composition of the Myriapoda in an Indonesian forest system, the fauna of the Halimun-Salak National Park in western Java was explored during the dry season (September-October 2015) in the framework of the German-Indonesian INDOBIOSYS project (Indonesian Biodiversity Discovery and Information System). A total of 980 Myriapoda specimens were collected by hand by 3-4 researchers from three different sites in the national park, from which 796 specimens were determined to a higher taxonomic level (class, order, family) and 617 specimens were determined to morphospecies. Among these, 27 were Symphyla (4%) (excluded from further analyses), 226 Chilopoda (28%) and 543 Diplopoda (68%). The Scolopendromorpha (64% of all identified centipedes) and Polydesmida (69% of all identified Diplopoda) were the most represented orders in our samples. Twenty-four morphospecies of Chilopoda were determined: one each of Scutigeromorpha and Lithobiomorpha, six Scolopendromorpha and sixteen Geophilomorpha. Nine orders of diplopods were present, with a total of 47 morphospecies: one each of Polyxenida, Glomeridesmida and Chordeumatida, two each of Glomerida, Spirobolida and Siphonophorida, seven of Sphaerotheriida, ten of Spirostreptida and 21 of Polydesmida. Two species curves were obtained to have a first idea about the myriapod diversity in the Halimun-Salak National Park and to compare the three individual collecting sites. Our results depict the Scolopendromorpha as the most common centipedes in Javanese rainforests and the Geophilomorpha as the most species-rich order. In contrast, the Polydesmida were the most dominant millipede group with 167 specimens and with 13 morphospecies the family Paradoxosomatidae was the most diverse.

A review of the family Trichopolydesmidae in North Africa with a description of a new species from Tunisia.

A new species of the genus Haplocookia Brölemann, 1915 is described from Cap Bon Peninsula in Tunisia (North Africa) and a historical account of the poorly understood genera Haplocookia and Heterocookia Silvestri, 1898 is provided. Comments on the taxonomy of the family Trichopolydesmidae are presented, as well as an identification key to the trichopolydesmid species hitherto known from North Africa and an updated list of the Polydesmida in the region.

Two centipedes new to the fauna of La Palma (Canary Islands, Spain) and notes on the Lithobiomorpha of the island (Chilopoda, Myriapoda).

Recent collecting on the island of La Palma has yielded two new records, Lithobius (Monotarsobius) crassipesoides and Lithobius (Lithobius) melanops, the former also being new to the Canary Islands and is recorded for the first time after its recent description. We additionally provide species records for the lithobiomorph species of the island, with new locality data and a distribution map based on recent and literature records. We update the checklist of the lithobiomorph species of the Canaries and document all the newly collected species.